Event related potentials (ERP) reveal a robust response to visual symmetry in unattended visual regions.

Visual symmetry at fixation generates a bilateral Event Related Potential (ERP) called the Sustained Posterior Negativity (SPN). Symmetry presented in the left visual hemifield generates a contralateral SPN over the right hemisphere and vice versa. The current study examined whether the contralateral SPN is modulated by the focus of spatial attention. On each trial there were two dot patterns, one to the left of fixation, and one to the right of fixation. A central arrow cue pointed to one of the patterns and participants discriminated its regularity (symmetry or random). We compared contralateral SPN amplitude generated by symmetry at attended and unattended spatial locations. While the response to attended symmetry was slightly enhanced, the response to unattended symmetry was still substantial. Although visual symmetry detection is a computational challenge, we conclude that the brain processes visual symmetry in unattended parts of the visual field.

The role of task on the human brain's responses to, and representation of, visual regularity defined by reflection and rotation.

Identifying and segmenting objects in an image is generally achieved effortlessly and is facilitated by the presence of symmetry: a principle of perceptual organisation used to interpret sensory inputs from the retina into meaningful representations. However, while imaging studies show evidence of symmetry selective responses across extrastriate visual areas in the human brain, whether symmetry is processed automatically is still under debate. We used functional Magnetic Resonance Imaging (fMRI) to study the response to and representation of two types of symmetry: reflection and rotation. Dot pattern stimuli were presented to 15 human participants (10 female) under stimulus-relevant (symmetry) and stimulus-irrelevant (luminance) task conditions. Our results show that symmetry-selective responses emerge from area V3 and extend throughout extrastriate visual areas. This response is largely maintained when participants engage in the stimulus irrelevant task, suggesting an automaticity to processing visual symmetry. Our multi-voxel pattern analysis (MVPA) results extend these findings by suggesting that not only spatial organisation of responses to symmetrical patterns can be distinguished from that of non-symmetrical (random) patterns, but also that representation of reflection and rotation symmetry can be differentiated in extrastriate and object-selective visual areas. Moreover, task demands did not affect the neural representation of the symmetry information. Intriguingly, our MVPA results show an interesting dissociation: representation of luminance (stimulus irrelevant feature) is maintained in visual cortex only when task relevant, while information of the spatial configuration of the stimuli is available across task conditions. This speaks in favour of the automaticity for processing perceptual organisation: extrastriate visual areas compute and represent global, spatial properties irrespective of the task at hand.

The extrastriate symmetry response is robust to alcohol intoxication.

Visual symmetry activates a network of regions in the extrastriate cortex and generates an event-related potential (ERP) called the sustained posterior negativity (SPN). Previous work has found that the SPN is robust to experimental manipulations of task, spatial attention, and memory load. In the current study, we investigated whether the SPN is also robust to alcohol-induced changes in mental state. A pilot experiment (N = 13) found that alcohol unexpectedly increased SPN amplitude. We followed this unexpected result with two new experiments on separate groups, using an alcohol challenge paradigm. One group completed an Oddball discrimination task (N = 26). Another group completed a Regularity discrimination task (N = 26). In both groups, participants consumed a medium dose of alcohol (0.65 g/kg body weight) and a placebo drink, in separate sessions. Alcohol reduced SPN amplitude in the Oddball task (contrary to the pilot results) but had no effect on SPN amplitude in the Regularity task. In contrast, the N1 wave was consistently dampened by alcohol in all experiments. Exploratory analysis indicated that the inconsistent effect of alcohol on SPN amplitude may be partly explained by individual differences in alcohol use. Alcohol reduced the SPN in light drinkers and increased it in heavier drinkers. Despite remaining questions, the results highlight the automaticity of symmetry processing. Symmetry still produces a large SPN response, even when participants are intoxicated, and even when symmetry is not task relevant.

The brain does not process horizontal reflection when attending to vertical reflection, and vice versa.

Previous work has found that feature attention can modulate electrophysiological responses to visual symmetry. In the current study, participants observed spatially overlapping clouds of black and white dots. They discriminated vertical symmetry from asymmetry in the target dots (e.g., black or white) and ignored the regularity of the distractor dots (e.g., white or black). We measured an electroencephalography component called the sustained posterior negativity (SPN), which is known to be generated by visual symmetry. There were five conditions with different combinations of target and distractor regularity. As well as replicating previous results, we found that an orthogonal axes of reflection in the distractor dots had no effect on SPN amplitude. We conclude that the visual system can processes reflectional symmetry in independent axis-orientation specific channels.

Right lateralized alpha desynchronization increases with the proportion of symmetry in the stimulus.

Research into the neural basis of symmetry perception has intensified in the last two decades; however, the functional role of neural oscillations remains unclear. In previous work Makin et al. (2014, Journal of Vision, 14, 1-12) and Wright et al. (2015, Psychophysiology, 52, 638-647) examined occipital alpha event-related desynchronization (alpha ERD). It was concluded that alpha ERD is right lateralized during active regularity discrimination but not during a secondary task. Furthermore, alpha ERD was unaffected by stimulus properties, such as the type of regularity. These conclusions are refuted by new time-frequency analysis on an electroencephalography (EEG) data set first introduced by Makin et al. (2020, Journal of Cognitive Neuroscience, 32, 353-366). We compared alpha ERD across five tasks. First, we found that right lateralization of alpha ERD was evident in all tasks, not just active regularity discrimination. This was caused by hemispheric differences in alpha power during prestimulus baseline (left < right), which equalized after stimulus onset (left = right). Second, we found that Alpha ERD increased with the proportion of symmetric elements in the image (PSYMM). Sensitivity to PSYMM was stronger on the right. These findings suggest that known extrastriate symmetry activations are accompanied by reduced alpha power.

Electrophysiological priming effects confirm that the extrastriate symmetry network is not gated by luminance polarity.

It is known that the extrastriate cortex is activated by visual symmetry. This activation generates an ERP component called the Sustained Posterior Negativity (SPN). SPN amplitude increases (i.e., becomes more negative) with repeated presentations. We exploited this SPN priming effect to test whether the extrastriate symmetry response is gated by element luminance polarity. On each trial, participants observed three stimuli (patterns of dots) in rapid succession (500 ms. with 200 ms. gaps). The patterns were either symmetrical or random. The dot elements were either black or white on a grey background. The triplet sequences either showed repeated luminance (black > black > black, or white > white > white) or changing luminance (black > white > black, or white > black > white). As predicted, SPN priming was comparable in repeated and changing luminance conditions. Therefore, symmetry with black elements is not processed independently from symmetry with white elements. Source waveform analysis confirmed that this priming happened within the extrastriate symmetry network. We conclude that the network pools information across luminance polarity channels.

The Formation of Symmetrical Gestalts Is Task-Independent, but Can Be Enhanced by Active Regularity Discrimination.

The brain can organize elements into perceptually meaningful gestalts. Visual symmetry is a useful tool to study gestalt formation, and we know that there are symmetry-sensitive regions in the extrastriate cortex. However, it is unclear whether symmetrical gestalt formation happens automatically, whatever the participant's current task is. Does the visual brain always organize and interpret the retinal image when possible, or only when necessary? To test this, we recorded an ERP called the sustained posterior negativity (SPN). SPN amplitude increases with the proportion of symmetry in symmetry + noise displays. We compared the SPN across five tasks with different cognitive and perceptual demands. Contrary to our predictions, the SPN was the same across four of the five tasks but selectively enhanced during active regularity discrimination. Furthermore, during regularity discrimination, the SPN was present on hit trials and false alarm trials but absent on miss and correct rejection trials. We conclude that gestalt formation is automatic and task-independent, although it occasionally fails on miss trials. However, it can be enhanced by attention to visual regularity.

Symmetric patterns with different luminance polarity (anti-symmetry) generate an automatic response in extrastriate cortex.

People can quickly detect bilateral reflection in an image. This is true when elements of the same luminance are matched on either side of the axis (symmetry) and when they have opposite luminance polarity (anti-symmetry). Using electroencephalography, we measured the well-established sustained posterior negativity (SPN) response to symmetry and anti-symmetry. In one task, participants judged the presence or absence of regularity (Regularity Discrimination Task). In another, they judged the presence or absence of rare colored oddball trials (Colored Oddball Task). Previous work has concluded that anti-symmetry is only detected indirectly, through serial visual search of element locations. This selective attention account predicts that the anti-symmetry SPN should be abolished in the Colored Oddball Task because there is no need to search for anti-symmetry. However, this prediction was not confirmed: The symmetry and anti-symmetry SPN waves were not modulated by task. We conclude that at least some forms of anti-symmetry can be extracted from the image automatically, in much the same way as symmetry. This is an important consideration for models of symmetry perception, which must be flexible enough to accommodate opposite luminance polarity, while also accounting for the fact anti-symmetry is often perceptually weaker than symmetry.

Electrophysiological responses to regularity show specificity to global form: The case of Glass patterns.

The holographic weight of evidence model (van der Helm & Leeuwenberg, J Math Psychol, 35, 1991, 151; van der Helm & Leeuwenberg, Psychol Rev, 103, 1996, 429) estimates that the perceptual goodness of moiré structures (Glass patterns), irrespective of their global form, is comparable to that of reflection symmetry. However, both behavioural and neuroscience evidences suggest that certain Glass forms (i.e. circular and radial structures) are perceptually more salient than others (i.e. translation structures) and may recruit different perceptual mechanisms. In this study, we tested whether brain responses for circular, radial and translation Glass patterns are comparable to the response for onefold bilateral reflection symmetry. We recorded an event-related potential (ERP), called the sustained posterior negativity (SPN), which has been shown to index perceptual goodness of a range of regularities. We found that circular and radial Glass patterns generated a comparable SPN amplitude to onefold reflection symmetry (in line with the prediction of the holographic model), starting approx. 180 ms after stimulus onset. Conversely, the SPN response to translation Glass patterns had a longer latency (approx. 400 ms). These results show that Glass patterns are a special case of visual regularity, and perceptual goodness may not be fully explained by the holographic identities that constitute it. Specialised processing mechanisms might exist in the regularity-sensitive extrastriate areas, which are tuned to global form configurations.

The extrastriate symmetry response can be elicited by flowers and landscapes as well as abstract shapes.

Previous research has investigated the neural response to visual symmetry. It is well established that symmetry activates a network of extrastriate visual regions, including V4 and the Lateral Occipital Complex. This symmetry response generates an event-related potential called the sustained posterior negativity (SPN). However, previous work has used abstract stimuli, typically dot patterns or shapes. We tested the generality of the SPN. We confirmed that the SPN wave was present and of similar amplitude for symmetrical shapes, flowers and landscapes, whether participants were responding either to image symmetry or to image color. We conclude that the extrastriate symmetry response can be generated by any two-dimensional image and is similar in different stimulus domains.

Representation of symmetry in the extrastriate visual cortex from temporal integration of parts: An EEG/ERP study.

When symmetry is present in the retinal image, a symmetry-sensitive network in the extrastriate visual areas activates, and response magnitude scales with degree of regularity. Is this activation driven by the regularity in the image, or can the network recover regularity of an object? We investigated whether the network responds to bilateral symmetry for dynamically occluded shapes, and thus responds to symmetry in the object. The stimulus was an irregular shape partly occluded by a rectangle. After 500 ms, the rectangle was displaced to the other side, occluding the previously visible half, and revealing the other half for 1000 ms. Therefore, no symmetry was present in the image at any point in time. Exp.1 and Exp.2 used vertical and horizontal axis of reflection, and in Exp. 3 there was no occluder. Participants could detect symmetry with >80% accuracy. More importantly, ERP analysis showed a symmetry-specific response from ∼300 ms after presentation of the second half of the shape. When integration was made from halves of asymmetric whole shapes (Exp.4), and when symmetry was not task-relevant (Exp.5), no symmetry response was recorded. The results show, for the first time, an electrophysiological evidence of symmetry representation in the brain obtained by assembling information over time into a unitary gestalt. The integration process occurs when observers look for symmetric matches between the parts, and only if these are perceived as belonging to the same object.

Visual perceptions of portion size normality and intended food consumption: A norm range model.

•Smaller portion sizes are associated with lower energy intake.•We test a norm range model of the portion size effect on intended intake.•A wide range of portion sizes were perceived as normal.•Portions perceived as normal did not prompt intended compensatory eating.•Portions perceived as smaller than normal prompted intended compensation.

Symmetry perception for patterns defined by color and luminance.

Perception of visual symmetry is fast and efficient and relies on both early low-level and late mid- and high-level neural mechanisms. To test for potential influences of early low-level mechanisms on symmetry perception, we used isoluminant, achromatic, and combined (color + luminance) patterns in a psychophysical and an event-related-potential (ERP) experiment. In the psychophysical experiment, pattern contrast was fixed at individual symmetry-discrimination threshold. Participants then judged whether a pattern was symmetric or random. Stimuli at isoluminance were associated with a large bias toward symmetry, achromatic stimuli introduced the opposite bias, and stimuli containing a balance of both color and luminance were perceived without bias. These findings are in line with distinct contrast sensitivity functions for color and luminance, with color providing low-frequency information useful for symmetry detection and luminance providing high-frequency information useful for detection of detail. The subsequent ERP experiment was run at high contrasts to assess processing of symmetry in suprathreshold conditions. Sustained posterior negativity, a symmetry-sensitive ERP component, was observed in all conditions and showed the expected dependence on symmetry. However, interactions between symmetry and contrast type were not observed. In conclusion, while our findings at threshold support models that propose an important contribution of low-level mechanisms to symmetry perception, at suprathreshold these low-level contributions do not persist. Therefore, under everyday viewing conditions, symmetry perception engages a relatively broad cortical network that is not constrained by low-level inputs.

An Electrophysiological Index of Perceptual Goodness.

A traditional line of work starting with the Gestalt school has shown that patterns vary in strength and salience; a difference in "Perceptual goodness." The Holographic weight of evidence model quantifies goodness of visual regularities. The key formula states that W = E/N, where E is number of holographic identities in a pattern and N is number of elements. We tested whether W predicts the amplitude of the neural response to regularity in an extrastriate symmetry-sensitive network. We recorded an Event Related Potential (ERP) generated by symmetry called the Sustained Posterior Negativity (SPN). First, we reanalyzed the published work and found that W explained most variance in SPN amplitude. Then in four new studies, we confirmed specific predictions of the holographic model regarding 1) the differential effects of numerosity on reflection and repetition, 2) the similarity between reflection and Glass patterns, 3) multiple symmetries, and 4) symmetry and anti-symmetry. In all cases, the holographic approach predicted SPN amplitude remarkably well; particularly in an early window around 300-400 ms post stimulus onset. Although the holographic model was not conceived as a model of neural processing, it captures many details of the brain response to symmetry.

Memory-guided tracking through physical space and feature space.

People can estimate the current position of an occluded moving target. This is called motion extrapolation, and it has been suggested that the performance in such tasks is mediated by the smooth-pursuit system. Experiment 1 contrasted a standard position extrapolation task with a novel number extrapolation task. In the position extrapolation task, participants saw a horizontally moving target become occluded, and then responded when they thought the target had reached the end of the occluder. Here the stimuli can be tracked with pursuit eye movements. In the number extrapolation task, participants saw a rapid countdown on the screen that disappeared before reaching zero. Participants responded when they thought the hidden counter would have reached zero. Although this stimulus cannot be tracked with the eyes, performance was comparable on both the tasks. The response times were also found to be correlated. Experiments 2 and 3 extended these findings, using extrapolation through color space as well as number space, while Experiment 4 found modest evidence for similarities between color and number extrapolation. Although more research is certainly needed, we propose that a common rate controller guides extrapolation through physical space and feature space. This functions like the velocity store module of the smooth-pursuit system, but with a broader function than previously envisaged.

Visual symmetry in objects and gaps.

It is known that perceptual organization modulates the salience of visual symmetry. Reflectional symmetry is more quickly detected when it is a property of a single object than when it is formed by a gap between two objects. Translational symmetry shows the reverse effect, being more quickly detected when it is a gap between objects. We investigated the neural correlates of this interaction. Electroencephalographic data was recorded from 40 participants who were presented with reflected and translated contours in one- or two-object displays. Half of the participants discriminated regularity, half distinguished number of objects. An event-related potential known as the Sustained Posterior Negativity (SPN) distinguished between reflection and translation. A similar ERP distinguished between one and two object presentations, but these waves summed with the SPN, rather than altering it. All stimuli produced desynchronization of 8-13 Hz alpha oscillations over the bilateral parietal cortex. In the Discriminate Regularity group, this effect was right lateralized. The SPN and alpha desynchronization index different stages of visual symmetry discrimination. However, neither component displayed the Regularity × Objecthood interaction that is observed in speeded discrimination tasks, suggesting that integration of visual regularity with objectness is not inevitable. Instead, both attributes may be processed in parallel and independently.

Examining visual complexity and its influence on perceived duration.

We investigated whether visual complexity of novel abstract patterns affects perceived duration. Previous research has reported that complex visual stimuli led to an underestimation of durations. However, to clarify the nature of the time estimation process, it is necessary to establish which component of image complexity, spatial or semantic, plays the critical role. Here we tested the impact of specific spatial properties. We used unfamiliar and abstract patterns made using black-and-white checkerboards in which the difference between stimuli was exclusively in configuration. Visual complexity was quantified by the GIF index based on a compression algorithm, which scanned the pattern in both horizontal and vertical directions. This metric correlated positively with subjective complexity (Experiment 1A). In the second study, we increased variability in the stimuli by changing the number of items across patterns while keeping overall size constant. A high positive correlation was found between objective and subjective complexity (r = 0.95) (Experiment 2A). In Experiments 1B and 2B, observers estimated pattern durations in seconds using a continuous scale. A multilevel linear analysis found that perceived duration was not predicted by visual complexity for either of the two sets of stimuli. These results provide new constraints to theories of time perception, hypothesizing that complexity leads to an underestimation of duration when it reduces attention to time.

When does perceptual organization happen?

Reflectional (mirror) symmetry is an important visual cue for perceptual organization. The brain processes symmetry rapidly and efficiently. Previous work suggests that symmetry activates the extrastriate cortex and generates an event related potential (ERP) called the Sustained Posterior Negativity (SPN). It has been claimed that no tasks completely block symmetry processing and abolish the SPN. We tested the limits of this claim with a series of eight new Electroencephalography (EEG) experiments (344 participants in total). All experiments used the same symmetrical or asymmetrical dot patterns. When participants attended to regularity in Experiment 1, there was a substantial SPN (Mean amplitude = -2.423 µV). The SPN was reduced, but not abolished, when participants discriminated dot luminance in Experiments 2 and 3 (-.835 and -1.410 µV) or the aspect ratio of a superimposed cross in Experiments 4 and 5 (-.722 and -.601 µV). The SPN also survived when the background pattern was potentially disruptive to the primary task in Experiment 6 (-1.358 µV) and when participants classified negative superimposed words in Experiment 7 (-.510 µV). Finally, the SPN remained when participants attended to the orientation of a diagonal line in Experiment 8 (-.589 µV). While task manipulations can turn down the extrastriate symmetry activation, they cannot render the system completely unresponsive. Permanent readiness to detect reflectional symmetry at the centre of the visual field could be an evolved adaptation.

Electrophysiological evidence of the amodal representation of symmetry in extrastriate areas.

Extrastriate visual areas are strongly activated by image symmetry. Less is known about symmetry representation at object-level rather than image-level. Here we investigated electrophysiological responses to symmetry, generated by amodal completion of partially-occluded polygon shapes. We used a similar paradigm in four experiments (N = 112). A fully-visible abstract shape (either symmetric or asymmetric) was presented for 250 ms (t0). A large rectangle covered it entirely for 250 ms (t1) and then moved to one side to reveal one half of the shape hidden behind (t2, 1000 ms). Note that at t2 no symmetry could be extracted from retinal image information. In half of the trials the shape was the same as previously presented, in the other trials it was replaced by a novel shape. Participants matched shapes similarity (Exp. 1 and Exp. 2), or their colour (Exp. 3) or the orientation of a triangle superimposed to the shapes (Exp. 4). The fully-visible shapes (t0-t1) elicited automatic symmetry-specific ERP responses in all experiments. Importantly, there was an exposure-dependent symmetry-response to the occluded shapes that were recognised as previously seen (t2). Exp. 2 and Exp.4 confirmed this second ERP (t2) did not reflect a reinforcement of a residual carry-over response from t0. We conclude that the extrastriate symmetry-network can achieve amodal representation of symmetry from occluded objects that have been previously experienced as wholes.

Lessons from a catalogue of 6674 brain recordings.

It is now possible for scientists to publicly catalogue all the data they have ever collected on one phenomenon. For a decade, we have been measuring a brain response to visual symmetry called the sustained posterior negativity (SPN). Here we report how we have made a total of 6674 individual SPNs from 2215 participants publicly available, along with data extraction and visualization tools (https://osf.io/2sncj/). We also report how re-analysis of the SPN catalogue has shed light on aspects of the scientific process, such as statistical power and publication bias, and revealed new scientific insights.